NEW YORK—Parked in front of a seductive portraitof Ann Coulter and an array of computer screens inside a home office in Harlem, Matt Braynard, former director of technology for Donald Trump’s presidential campaign, welled up with emotion as he recounted his experience attending Trump’s inauguration a week earlier.

It was not the ceremony itself, but what happened afterward, when Braynard spotted a pale teenager in a red hat and a gray hoodie standing on a street corner near the Capitol holding a sign with a picture of a cartoon frog and the message “LOVE NOT HATE.” Struck by the display, Braynard made a beeline for the kid, shook his hand and enlisted his wife to take a photograph of them together.

“I felt like I had found a friend that I hadn’t spoken to in a long time,” recalled Trump’s misty-eyed onetime data guru, who had never met the teenager before in his life. “It was, like, immediate connection.”

The chance encounter was a rare in-real-life meeting of veterans of the Great Meme War. That’s the grandiose name given—only half ironically—to the decentralized efforts of a swarm of anonymous internet nerds to harass Trump’s detractors and flood the Web with pro-Trump, anti-Hillary Clinton propaganda. Their weapons of choice were memes, bits of reproducible culture whose most recognizable form is shareable internet photos, like cats behaving adorably or Clinton sending a text message, with captions meant to be funny.

Braynard showed me a badge he ordered online to memorialize his service in the Great Meme War. It features Pepe the Frog—a cartoon symbol of mischievous fun or racist hatred, depending on whom you ask—and the name of his make-believe battalion, “The 1st Deplorables.”

Veterans of the Great Meme War brag that they won the election for Trump. Just about everyone else, if they’re aware of these efforts at all, assumes they amounted to little more than entertainment for bored geeks and some unpleasant episodes for the targets of its often racist and sexist harassment campaigns. After all, the idea that a swarm of socially alienated trolls played a meaningful role in a multibillion-dollar presidential campaign by, among other gambits, relentlessly spreading images of a cartoon frog is at least as ridiculous as the idea that a billionaire TV entertainer could win that campaign.

There is no real evidence that memes won the election, but there is little question they changed its tone, especially in the fast-moving and influential currents of social media. The meme battalions created a mass of pro-Trump iconography as powerful as the Obama “Hope” poster and far more adaptable; they relentlessly drew attention to the tawdriest and most sensational accusations against Clinton, forcing mainstream media outlets to address topics—like conspiracy theories about Clinton’s health—that they would otherwise ignore. And they provoked a variety of real-world reactions, from Clinton’s August speech denouncing the alt-right to the Anti-Defamation League’s designation of Pepe as a hate symbol to—after the election—the armed assault on a Washington pizzeria wrongly believed to be hiding sex slaves.

Part of the power of memes has always been their organic, grass-roots quality: They bubble up from the fever swamps of the internet, shrouded in anonymity, as agents of chaos and mockery. But in this election, something seemed to change. They began colliding with a real campaign operation and doing useful work, seemingly always pushing in one direction. Curious about what happened, I tracked down and interviewed a number of veterans of the Great Meme War, along with others who hung out in the same dark corners of the internet and watched it all unfold. It turns out that, as anonymous online pranksters go, they’re surprisingly organized and motivated. It also turns out that the Trump campaign, which spent relatively little on messaging, paid rapt attention to meme culture from the start. It took it seriously, even pushing some memes out to the candidate’s millions of Twitter followers.

Trump’s campaign will not be the last to tap into this subculture. Internet troll Charles Johnson, a self-commissioned general in the Great Meme War with close ties to Trump’s political operation, claimed he has fielded about a dozen post-election phone calls from the Washington area about the political potential of memes. “If you’re trying to win an election and you have a million dollars to spend on political ads or $100,000 to spend on trolling,” he said, “I would advise everyone to spend the hundred thousand on the troll.”

If the soldiers in the Great Meme War are even partly right about their capabilities, then their efforts have profound implications for the future of politics. But before tackling that question, it is worth asking how, in the first place, a community of some of the savviest, most subversive internet users became a hotbed of support for a 70-year-old white billionaire who refers to Apple products as “damn computers and things.” And for that matter, what exactly is a meme, anyway?

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The concept of a “meme,” in its broadest sense, has been around for decades. The term was first coined in 1976 by evolutionary biologist Richard Dawkins, who defined a meme as “a unit of cultural transmission or a unit of imitation”—essentially, a reproducible bit of the DNA of human culture. He saw the idea expansively; the most effective memes, like religious rituals and catchy melodies, worm their way into people’s brains, spreading across entire societies and shaping human behavior for generations.

The Jesus fish is an ancient meme, and Uncle Sam is an early American meme. The planking fad, in which people lie flat on their fronts in weird places and pose for photographs, is a recent behavioral meme. The term came into popular parlance with the advent of the “internet meme,” usually a photograph with a clever caption that is shared around the Web. Created anonymously, remixed endlessly and shared constantly, the most viral memes seem to materialize out of nowhere.

But the typical internet meme doesn’t exactly come from nowhere. Its very Darwinian life cycle often begins among thousands of other memes on a group of obscure message boards frequented by the internet’s most devoted users, mostly young men, who Photoshop captioned images for their own amusement. The most promising become popular on these boards, as users post their own variations on the theme, and end up crossing over to more mainstream platforms like Reddit and Tumblr, which are used by “normies,” or normal people, and often drive what’s popular on the internet at any given time. From there, the most successful memes start populating platforms that almost everyone uses, like Facebook, and a very select few, like LOLCats and Rickrolling,enter the cultural canon, becoming recognizable even to one’s parents.

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The fighters in the Great Meme War took their intimate knowledge of this ecosystem and weaponized it, genetically engineering pro-Trump and anti-Clinton supermemes they designed to gain as much mainstream traction as possible. They juiced the rules on platforms like Reddit and created networks of fake accounts on Twitter to push the memes in front of as many eyeballs as possible as quickly as possible. The staging ground was an anonymous message board called “/pol/”—the “politically incorrect” section of 4Chan, which was founded in 2003 to host discussions about anime and has since evolved into a malignant hive mind with vast influence over online culture. The denizens of /pol/ believe that their efforts memed President Trump into existence, midwifing his presidency from a far-fetched fantasy into our current reality. Memes like “the Trump Train” were popularized by 4Chan, spread to the rest of the Web and then rapidly absorbed into official campaign messaging—sometimes reaching all the way to the candidate himself.

Braynard, who led Trump’s data team from October 2015 through March 2016, said younger staffers would regularly pass around memes as morale boosters. He cited a video called “You can’t stump the Trump,” a phrase first popularized on 4Chan, that mashed up Trump’s early primary debate highlights with a narrator from a nature documentary talking about centipedes and other campy effects, as a particular favorite of staffers. In October 2015, months before Pepe became a subject of campaign controversy, Trump tweeted a link to the video along with an illustration of the cartoon frog with Trump’s hair standing in front of the presidential seal. The clip has been viewed more than a million times.

But it wasn’t until the arrival months later of Steve Bannon, who brought with him an in-depth knowledge of the internet’s underbelly acquired while growing the anti-establishment Breitbart News, that the campaign’s engagement with the fever swamps reached its apogee. By the fall, a team in the war room at Trump Tower was monitoring social media trends, including The_Donald subreddit—a message board that acted as a conduit between 4Chan and the mainstream Web and refers to its users as “centipedes” in honor of the aforementioned video—and privately communicating with the most active users to seed new trends, according to two former Trump campaign officials. The team would bump up anything particularly catchy to social media director Dan Scavino. (Trump spokeswoman Hope Hicks said there were no staffers dedicated full-time to monitoring social media trends, and that Bannon was not involved in social media strategy.) But one former campaign official said the goal was to relentlessly tilt the prevailing sentiment on social media in favor of Trump: “He clearly won that war against Hillary Clinton day after day after day.”

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People who spend their days on internet message boards tend not to be political insiders, and even by internet standards 4Chan has always had an outsider bent, which might explain why it went so hard for a burn-it-all-down outsider like Trump. During the early 2000s, the boards were vehemently anti-George W. Bush, a hub for 9/11 truthers as well as for trolling religious conservatives. The vaguely leftist hacking collective Anonymous grew out of the Bush-era boards. Later, 4Chan fell for Ron Paul, helping fuel the quirky Texas congressman’s unlikely internet stardom. The boards also developed a culture of hard-core racist language; at first, this gratuitous bigotry was motivated primarily by a desire to get a rise out of normies, but it fostered an environment in which genuine racists felt at home, too.

In 2011, 4Chan created /pol/, its politically incorrect board, in part to house racist threads and other rants that were polluting the rest of the site. The white nationalist alt-right was forged in the crucible of 4Chan and remains indelibly marked by its emergence from meme culture. Screenshots from hacked social media accounts belonging to the late Trayvon Martin first surfaced on /pol/ as part of a campaign of posthumous character assassination that painted the African-American teenager as a pot-smoking delinquent who had it coming.

The site has also grown increasingly preoccupied with gender politics. “If you’re young, white and male, you’re on 4Chan,” says Johnson, who recently tried to buy the website. That’s clearly an exaggeration, but 4Chan has demonstrated an alarming power to whip up misogyny against its perceived enemies. The “Gamergate” harassment campaigns against women in the video-game industry were often organized on 4Chan. That episode involved allegations that a female video-game developer was cheating on her boyfriend with multiple men in the industry, and thus ushered the term “cuck,” short for cuckold, into 4Chan parlance. (This, in turn, had profound linguistic consequences for the Republican presidential primary when “cuckservative” became the insult of choice for insufficiently Trumpian Republicans.) Gamergate also hardened anti-political correctness sentiment on 4Chan, and when administrators eventually banned discussion of the topic, it proved a boon to the nascent 8Chan—a sort of ISIS to 4Chan’s Al Qaeda, a splinter group whose founder believed 4Chan had grown too controlled.

These new strains festered on the boards until June 16, 2015, when beauty pageant owner and former Pizza Hut pitchman Donald Trump arrived on the scene like a gift from the troll gods. Announcing his presidential candidacy, Trump disparaged Mexican immigrants in comically insulting language as paid actors cheered him on. The Chans went wild.

“For a lot of people, on the first day it was like, ‘This would be fucking hilarious,’ and then when he started coming up with policy stuff—the border wall, the Muslim ban—people on the boards were like, ‘This can’t be real. This is the greatest troll of all time,’” recounted “Marcus,” a former military intelligence officer in his 30s who participated in the Great Meme War. Marcus, who lives in Washington state, professionally monitors message boards across the Web and spoke on condition of anonymity. (He initially arranged to speak to me through a mutual acquaintance who gave me a burner number with a Rhode Island area code and an appointment set in Mountain Time.)

A thread posted on /pol/ the day after Trump’s announcement captured the essence of the board’s sentiment for the rest of the primaries. Titled “Jeb Bush the cuck,” it lists a number of knocks against the then-GOP front-runner, including “no jawline,” “cucked by every white woman so had to marry a Mexican” and “Makes Mexican babies.” The first response under the original post is just a picture of Trump with the caption, “You’re fired.”

As Trump’s list of apostasies grew and his candidacy continued to soar, the tone of the Chans’ support shifted perceptibly, says Marcus. “It went from ironic to militant very quickly.” His observation speaks to the enigmatic and ephemeral nature of the boards, which are anonymous and automatically delete most threads after a few hours or days. No one knows whether 4Chan’s Ron Paul diehards of yesteryear have become the Trump fanatics of today, or whether the boards are now just drawing a different crowd.

Without a doubt, many participants are genuine Trump supporters. A person close to the Trump campaign introduced me to “Daniel,” a young man who professed to have friends in the White House. A frequent /pol/ and 8Chan poster, he told me he created several fake personas on Reddit and one on Twitter to post anti-Clinton agitprop. “The reason I fought in the meme war is that as Andrew Breitbart said we are at literal war with the left. There is an ideological Cold War going on right now and the victor will determine the fate of Western Civilization,” Daniel wrote in an email.

Other meme warriors simply think there is no greater cosmic joke than electing Trump president. “Most of the people who took part in the Great Meme War hate Trump a lot,” insisted Gregg Housh, a reformed hacker and active 4Chan user who did a stint in federal prison a decade ago and was an early ringleader of Anonymous. (Linking his real-world name to his online activity makes him a “namefag” in the eyes of /pol/, which is populated mostly by “anonfags.” In the world of /pol/, everyone is some sort of fag.)

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The problem is compounded by the fact that deception and trolling are at the heart of 4Chan’s culture, making it impossible to strike through the pasteboard mask of anonymity and determine a poster’s true motivation. Even for those who study 4Chan professionally, the boards remain a hall of mirrors.

“We don’t know how much was sincere, and the outcome of the election does not answer the question either,” said Whitney Philips, a professor of literary studies at Mercer University in Georgia whose research focuses on trolling and other internet subcultures. “We don’t know before the election how much of the support was people messing around.”

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Real or not, 4Chan’s enthusiasms have a habit of making their way to larger, more powerful platforms. Take the “cuck” trope: In July 2015, a /pol/ user helpfully linked to a blog post on an alt-right website that explained, “A cuckservative is a white (non-Jewish) conservative who isn’t racially aware.” A week later, on July 22, Rush Limbaugh introduced the term to his 13 million listeners when he called Trump’s critics within the party “cuckolded Republicans.” The political usage of “cuck” quickly exploded both on the boards and more mainstream channels, like Breitbart, whose former tech editor Milo Yiannopoulos described it as a “gloriously effective insult” and denied the term’s current usage carried racial connotations.

Through Yiannopoulos, who before his resignation in February often authored laudatory posts about 4Chan, the sexagenarian Bannon gained an appreciation for meme subculture, and Breitbart came to embrace it. “When Bannon said [Breitbart] was the platform for the alt-right, this is what he was talking about,” said a person who worked with Bannon at Breitbart. “He didn’t mean Richard Spencer. He meant the trolls on Reddit or 4Chan.”

In May, Breitbart published an attack on its ex-columnist Ben Shapiro authored by “Pizza Party Ben,” a pseudonymous participant in the Great Meme War who has appeared onstage for Yiannopoulos’ campus tour as a “meme-ology” consultant. (Pizza Party Ben declined an interview request, calling me “a fag.”) Yiannopoulos also recruited Anthime Gionet, a former BuzzFeed social media strategist who goes by the online nom de guerre Baked Alaska, to do work for Breitbart. Gionet, an active pro-Trump meme warrior, told me that in his free time he summoned “vast meme armies” for Trump on 4Chan and the website 9GAG.

Under Bannon, Breitbart mastered the art of the viral image to further its own brand. A recent Columbia Journalism Reviewanalysis found that while images constituted only 5 percent of the posts on Breitbart’s Facebook page last year, they accounted for half of the page’s 100 most-shared posts.

Those lessons infused the Trump team’s approach to sites like 4Chan, but the campaign would keep its distance, rather than trying to engage directly with their notoriously fickle, hostile communities. “They were more afraid of that than they were excited about how to use it,” said a former Trump campaign staffer. “This is a community that can, like, flip really easily and has the ability to either love or hate based on a single post. They had to be very careful about what they were doing.”

But its use of memes landed the Trump campaign in some pretty unsavory company. In November 2015, Trump tweeted a virulently racist image titled “USA CRIME STATISTICS ~ 2015” that depicted a menacing black man holding a gun alongside made-up statistics overstating the proportion of murders committed by black Americans. While the source of the chart was traced to a neo-Nazi Twitter account, the image of the gangbanger had been floating around on 4Chan for some time. In July, Trump’s campaign tweeted a picture that had been circulating on 8Chan that superimposed Hillary Clinton’s face on a background of $100 bills with the caption “Most Corrupt Candidate Ever” written on a six-sided star that, given the boards’ anti-Semitic proclivities, was almost certainly a Star of David. (The Trump campaign insisted unconvincingly that it was a sheriff’s star.)

Democrats dabbled with memes—the Clinton campaign even built a meme generator that was quickly swarmed by pro-Trump trolls—but they were generally dismissive of these efforts. A former Clinton aide, who spoke on condition of anonymity due to the requirements of a new job, scoffed at the notion that memes played a meaningful role in the campaign. “If you see a Nazi frog glorifying Donald Trump and you’re on the fence and you’re like, ‘OK, now I’m going to vote for Donald Trump,’ you were never going to vote for Hillary Clinton anyways,” said the aide. But the aide conceded that such efforts could have discouraged uncommitted voters from going to the polls, and others on the left view the meme ecosystem as a real asset to Trump.

“Those message boards matter,” said Angelo Carusone, president of Media Matters for America, an advocacy group founded by Clinton ally David Brock. Carusone pointed to the boards’ role in pushing conspiracy theories about Clinton’s health and their lesser-known work trying to discredit Clinton in the eyes of black voters, in part with memes invoking her use of the racially tinged term “superpredator” in 1996. Carusone also said that the boards’ organized harassment across the internet—“raids” of other social media platforms and comment sections—discouraged expressions of pro-Clinton sentiment from people caught off guard by the vitriol. The former Clinton aide said such harassment contributed to a tendency among Clinton supporters to congregate in secret, members-only Facebook groups, where they were preaching to the proverbial choir.

Meanwhile, 4Chan users and the alt-right movement it spawned drew increasing attention from prestigious corners of “normie” society, achieving a central goal of their trolling. In August, Clinton delivered a speech devoted to condemning the alt-right, which the boards took as a testament to “meme magic,” the ability of their trolling to create real-world effects. In September, the Anti-Defamation League classified Pepe the Frog as a hate symbol, another coup for Trump’s white nationalist fans, who had been busy bedecking the frog with swastikas and other racist trappings.

Then, in October, WikiLeaks began releasing hacked emails from the account of Clinton campaign chairman John Podesta, setting up the most disturbing meme magic yet. What many of the boards’ most active users lack in social graces they make up for in the ability to process and find patterns in high volumes of information, a talent they only half-jokingly call “weaponized autism.” The 4Chan hive-mind was made for the massive WikiLeaks dump, which brought new releases nearly daily for the final month of the campaign, and the boards frantically combed through them.

In the final days of the campaign, activity on the boards reached a fever pitch, and their users struck meme gold with the release five days before the election of a 2015 email in which Podesta’s brother Tony, a prominent lobbyist, invited him to attend a “spirit cooking” session in New York with the Serbian performance artist Marina Abromovic. A fixture in chi-chi Manhattan circles, her 1996 Spirit Cooking cookbook calls for such ingredients as breast milk, semen and “13,000 grams of jealousy.” Abromovic’s appearance in Podesta’s emails was a godsend for a community desperate to portray Clinton as a literal witch.

“People were pushing Spirit Cooking all over Spanish-language and African-American social media channels. A lot of people have sock-puppet accounts and they pretend to be black. And it worked. It got traction everywhere,” recounted Marcus. “I know tons of people who were like, ‘My abuela, as soon as she saw this, said, “If you vote for Hillary Clinton I’ll disown you.”’”

With Spirit Cooking popping on social media, the scene was set for an even more deranged attack on Clinton: Pizzagate. On November 2, a /pol/ user authored a post claiming that references to pizza in Podesta’s emails were code for pedophilia. 4Chan users were primed to make this bizarre (and completely unsubstantiated) connection, Marcus explained, because the site has a long history of linking to child pornography and because users long ago began shortening child porn to “CP” to evade detection, which had evolved into the code word “cheese pizza.”

Like much else on /pol/, it is unclear to what extent posters were trolling and to what extent they genuinely believed they were sleuthing out a child sex ring. “Let’s meme this into reality, it’s too good,” wrote one user on one of the original Pizzagate threads. “It was absolutely a joke and a guy just made it up on the spot,” Housh said. “I was on the thread and people thought it was hilarious and halfway through they were like, ‘How can we get people to take this seriously?’”

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Take it seriously people did.Pizzagate quickly supplanted Spirit Cooking as the boards’ closing argument. /Pol/ users combed through Podesta’s emails for other references to pizza and developed an elaborate conspiracy theory positing a Clinton-linked sex ring run out of a D.C. pizzeria owned by Brock’s ex-boyfriend. They created memes and charts and pushed them to broader audiences on The_Donald subreddit. Twitter users with big followings like blogger Mike Cernovich and Pizza Party Ben began tweeting about the theory with the hashtag #PizzaGate.

Trump won the election, and the Chans and The_Donald took their victory laps. But Pizzagate continued to fester, and in December, an impressionable North Carolina man heard about the fake sex ring and began researching it online. Carrying an assault rifle, he stormed the D.C. pizzeria that he believed housed the sex slave ring. Finding only pizza, he surrendered to the police. (“I regret how I handled the situation,” the man told the New York Times from jail.) One other Pizzagate casualty was Michael Flynn Jr., the son of Trump’s since-ousted national security adviser who was fired from the Trump transition team for his role in spreading the bogus story.

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In the wake of Trump’s win, interest in meme warfare is only growing. NATO has begun studying the potential role of memes in its struggle against the Islamic State. In January, the Kremlin tweeted an image of Pepe from the account of its embassy in London in a nod to the broadly pro-Trump, pro-Russia, nationalist coalition that has adopted Pepe as its calling card.

Meanwhile, the meme warriors have opened up a second front in Europe, where they are applying their U.S. election experiences to help far-right parties in upcoming races in France and Germany. “We’ve built a whole team in France. We’re in the process of building one in Germany,” the white nationalist hacker Andrew Auernheimer, aka Weev, a 4Chan veteran, told me a month after Trump’s win. “We’re about to get back in the saddle. Start making trouble again.”

Auernheimer, who once worked in marketing, said he and a team of unnamed accomplices had used industry-grade marketing tools like multivariate testing to deploy the most effective pro-Trump memes online during the U.S. election. He said he studied persuasion literature and evangelical street preaching to perfect his methods of converting Bernie Sanders supporters into Trump backers, and that he was in the process of recruiting volunteer teams in Germany and France to tip those countries to nationalist parties.

He is far from alone in targeting Europe.In January, BuzzFeed gained access to private chat rooms in which veterans of the Great Meme War were organizing to influence the French elections. The highly structured campaign included long guides that provided prefabricated memes and instructions for English speakers who want to appear French on social media. Organizers are urging American users to set up fake “sock-puppet” social media accounts and to teach French collaborators how to create effective memes in an effort to help the far right’s anti-immigrant, pro-Russia candidate, Marine Le Pen.

In our call, Auernheimer, breaking at points into manic laughter, pointed to growing friction between France’s military and police and its civilian leadership, and said he was working to foment popular sentiment in favor of a coup if Le Pen’s National Front does not win in April. “We hope to catalyze a new democratic system or encourage military and police putting the country back in order,” he said.

He offered to show me his command center if I flew out to his home in Ukraine. In January, when I emailed him to follow up on his offer, he responded, “Due to an NDA I can’t have any further discussions with you on this subject. Sorry.” It is not clear whether the non-disclosure agreement, like meme magic itself, is real or illusory. We may find out in April.